THE CHLOROPHYLL-CARBON DIOXIDE RATIO DURING PHOTOSYNTHESIS

Using a rapid spectrographic method of carbon dioxide measurement previously described by McAlister (1937) further studies on the time course of photosynthesis in the higher plant, wheat, variety Marquis, are herein reported. Of major importance in this work is the discovery of a pick-up of carbon dioxide in darkness immediately following a high rate of photosynthesis (see Figs. 3 and 4). This pick-up is believed to be due to the action of a carbon dioxide-combining intermediate; i.e., the "acceptor molecule" for carbon dioxide in photosynthesis. The conditions under which this phenomenon has so far been observed indicate that the intermediate is formed in relatively large quantities during the actual process of photosynthesis and not before. That the intermediate is chlorophyllous in nature is suggested by a simple stoichiometry of the order of unity that is found to exist between the number of carbon dioxide molecules taken up and the total number of chlorophyll molecules present in the plant. This is in opposition to the idea of a large photosynthetic unit of some 2000 chlorophyll molecules operating together in the reduction of 1 carbon dioxide molecule. Further studies of the induction phase under various conditions of previous dark rest and of carbon dioxide and light limitation are herein described. Employing the simple hypothesis that the number of carbon dioxide molecules not reduced during the induction period (induction loss) gives a measure of the number of elementary photosynthetic cycles unoperative or compensated for during induction together with the experimental fact that this induction loss is of the order of the total number of chlorophyll molecules present, these latter studies also indicate, in a less direct manner, that chlorophyll participates in photosynthesis as an individual molecule and not as part of a very large multimolecular chlorophyll unit. The fast dark reaction lasting about 1 minute (Fig. 7) required to reproduce both (a) the phenomena of induction in carbon dioxide assimilation and (b) the recovery of fluorescence of chlorophyll in leaves in darkness as observed by Franck and Wood (1936), demonstrates a close relationship between the fluorescence of chlorophyll and induction in photosynthesis. The rate of respiration (carbon dioxide production) of the higher plant, wheat, was measured under intense illumination and in the absence of carbon dioxide (to suppress assimilation). This value was found to be identical with the dark respirational rate measured before and after the light period, indicating very positively the absence of any direct effect of light on respiration.     

REDUCTION OF CARBON DIOXIDE COUPLED WITH THE OXYHYDROGEN REACTION IN ALGAE

1. Unicellular algae possessing a hydrogenase system (Scenedesmus and other species), and having been adapted by anaerobic incubation to the hydrogen metabolism, reduce oxygen to water according to the equation O₂ + 2H₂ → 2H₂O. 2. The oxyhydrogen reaction proceeds undisturbed only in the presence of carbon dioxide, which simultaneously is reduced according to the equation CO₂ + 2H₂ → H₂O + (CH₂O) = (carbohydrate). 3. The maximum yield of the induced reduction is one-half molecule of carbon dioxide reduced for each molecule of oxygen absorbed. 4. Partial reactions are recognizable in the course of the formation of water and it is with the absorption of the second equivalent of hydrogen that the carbon dioxide reduction appears to be coupled. 5. The velocity of the reaction increases in proportion to the partial pressure of oxygen, but only up to a certain point where any excess of oxygen causes the inactivation of the hydrogenase system. The reaction then ends prematurely. 6. During the oxyhydrogen reaction little or no oxygen is consumed for normal respiratory processes. 7. Small concentrations of cyanide, affecting neither photosynthesis nor photoreduction in the same cells, first inhibit the induced reduction of carbon dioxide and then lead to a complete inactivation of the hydrogenase system. 8. Hydroxylamine, added after adaptation, has either no inhibitory effect at all, or prevents solely the induced reduction of carbon dioxide without inactivating the hydrogenase system. 9. Dinitrophenol prevents the dark reduction of carbon dioxide while the reduction of oxygen continues to the formation of water. 10. Glucose diminishes the absorption of hydrogen, probably in its capacity as a competing hydrogen donor. 11. The induced reduction of carbon dioxide can be described as an oxido-reduction similar to that produced photochemically in the same cells.     

Measurement of Distribution of Photosynthesis in Plant Canopies

ATTEMPTS have been made to measure or calculate photosyn-thetic activity in various morphological parts or height layers of plant canopies. This has been done by measuring dry weight changes in relation to removal of the parts¹, carbon dioxide exchange when the parts were enclosed in assimilation chambers² and a combination of progressive defoliation and whole canopy carbon dioxide uptake measurement^3,4. To study the activity in layers of the canopy, Leach and Watson⁵ placed phytometers at various points in the canopy to stimulate the photosynthesis of adjacent plant parts and aerodynamic methods have since been used to estimate the distribution of photosynthesis in canopies⁶. By using photosynthesis response data for leaves and information about the environment in the profile, Monsi and Saeki⁷ and others^8,9 have estimated localized photosynthesis.     

Carbon dioxide assimilation from air and water by duckweed Spirodela polyrrhiza (L.) Schleid

Carbon dioxide assimilation by duckweed, S. polyrrhiza, was measured using a glass assimilation box and ¹⁴C-NaHCO₃, under different pH conditions of water. S. polyrrhiza assimilates carbon dioxide from both air and water. The carbon assimilation from air is comparable to the assimilation from water under normal pH conditions.     

OXIDATIVE ASSIMILATION IN RELATION TO PHOTOSYNTHESIS IN CHLORELLA

1. An oxidative assimilation of acetic add and glucose in darkness has been demonstrated in the green alga, Chlorella pyrenoidosa. From manometric experiments it has been shown that 1 mol (CH₂O) per mol acetic add and 5 mols (CH₂O) per mol glucose are produced. 2. The time required for complete utilization of a limited amount of acetic acid or glucose is not affected by illumination in the absence of carbon dioxide. 3. The time required for complete utilization of a limited amount of glucose is not affected by the simultaneous occurrence of photosynthesis. It must therefore be concluded that the accumulating product of photosynthesis cannot be glucose but must be some slowly respirable (storage) material. 4. Possible interrelationships between oxidative assimilation and photosynthesis may be further studied by following, in darkness and in light, the time course of oxidative assimilation of substrates which are possible intermediates in the two processes.     

Root respiration associated with ammonium and nitrate absorption and assimilation by barley

We examined nitrate assimilation and root gas fluxes in a wild-type barley (Hordeum vulgare L. cv Steptoe), a mutant (nar1a) deficient in NADH nitrate reductase, and a mutant (nar1a;nar7w) deficient in both NADH and NAD(P)H nitrate reductases. Estimates of in vivo nitrate assimilation from excised roots and whole plants indicated that the nar1a mutation influences assimilation only in the shoot and that exposure to NO₃⁻ induced shoot nitrate reduction more slowly than root nitrate reduction in all three genotypes. When plants that had been deprived of nitrogen for several days were exposed to ammonium, root carbon dioxide evolution and oxygen consumption increased markedly, but respiratory quotient—the ratio of carbon dioxide evolved to oxygen consumed—did not change. A shift from ammonium to nitrate nutrition stimulated root carbon dioxide evolution slightly and inhibited oxygen consumption in the wild type and nar1a mutant, but had negligible effects on root gas fluxes in the nar1a;nar7w mutant. These results indicate that, under NH₄⁺ nutrition, 14% of root carbon catabolism is coupled to NH₄⁺ absorption and assimilation and that, under NO₃⁻ nutrition, 5% of root carbon catabolism is coupled to NO₃⁻ absorption, 15% to NO₃⁻ assimilation, and 3% to NH₄⁺ assimilation. The additional energy requirements of NO₃⁻ assimilation appear to diminish root mitochondrial electron transport. Thus, the energy requirements of NH₄⁺ and NO₃⁻ absorption and assimilation constitute a significant portion of root respiration.     

The Gas Exchange of Hydrogen-adapted Algae as Followed by Mass Spectrometry

A mass spectrometer inlet and an oxygen electrode in the same vessel allowed the continuous recording of the gases exchanged (H₂, CO₂, O₂) by hydrogenase-containing anaerobically adapted Scenedesmus obliquus strain D₃ (Gaffron) and Chlorella fusca Shihira et Krauss (= pyrenoidosa) 211-15. A light intensity which produces more photosynthetic oxygen than the cells can re-reduce to water leads to de-adaptation and the substitution of normal photosynthesis for photoreduction. The sequence of these metabolic events was recorded in a matter of a few minutes. Upon exposure of these adapted algae to light, an evolution of hydrogen lasting up to 60 seconds preceded any other light-dependent gas exchange. In the presence of 3-(3,4-dichlorophenyl)-1,1-dimethylurea, this initial hydrogen production was inhibited approximately 50%, pointing to a contribution of electrons by photosystem II. At very low hydrogen tensions (0.1 microliter per milliliter), a balance between light-induced production and absorption of hydrogen was observed in normal, unpoisoned algae. Addition of either glucose or inhibitors of phosphorylation increased the release of hydrogen in the light very considerably. When the light was turned off the algae consumed the remaining amount of hydrogen, only to release it again upon illumination. This reversible hydrogen exchange persisted even when any concomitant carbon dioxide exchange had been abolished.     

Carbon dioxide exchange in a high-arctic fen estimated by eddy covariance measurements and modelling

The high-arctic environment is an environment where the consequences of global warming may be significant. In this paper we report on findings on carbon dioxide and water vapour fluxes above a sedge-dominated fen at Zackenberg (74°28′N, 20°34′ W) in The National Park of North and East Greenland. Eddy covariance measurements were initiated at the start of the growing season and terminated shortly before its end lasting 45 days. The net CO₂ flux during daytime reaches a high of 10 μmol m^–2s^–1, and around the summer solstice, net CO₂ assimilation occurred at midnight, resulting in net carbon gain during the night. The measured carbon dioxide fluxes compare well to estimates based on the photosynthesis model by Collatz et al. (1991 ). The total growing-season net ecosystem CO₂ exchange was estimated to be 96 g C m^–2 based on the carbon dioxide model and micrometeorological data. Finally, the combined CO₂ assimilation and soil respiration models are used for examining the dependence of the carbon dioxide budget on temperature. The ecosystem is found to function optimally given the present temperature conditions whereas either an increase or a decrease in temperature would reduce the ecosystem CO₂ accumulation. An increase in temperature by 5 °C would turn the ecosystem into a carbon dioxide source.     

Carbon dioxide exchange in the needles of the common spruce in southern taiga spruce forests

Dynamics of carbon dioxide exchange in the Common Spruce (Picea abies L.) in relation to environmental factors was monitored during several seasons. Direct linear dependence of photosynthesis rate from the levels of air temperature and illumination was found, and correlation coefficients were 0.860 (p < 0.001) and 0.704 (p < 0.001). It was found that seasonal maximum of net photosynthesis production was attained at temperatures of 23–25°C. A decrease in temperature optimum was associated with reduction of the CO₂ assimilation intensity level. The impact of environmental factors on photosynthesis intensity is discussed in terms of the developed model. Using this model, we demonstrated that temperature and illumination dynamics in toto accounts for 82% of changes in photosynthesis rate. It is the air temperature that exerts the strongest influence on the process of photosynthesis. According to our calculations, the net photosynthesis level was three times higher than the level of respiration. This is indicative of a positive carbon dioxide balance in the needles of the Common Spruce.     

SOME EFFECTS OF DERIVATIVES OF VITAMIN K ON THE METABOLISM OF UNICELLULAR ALGAE

Vitamin K₁, 2-methyl-3-phytyl-1,4-naphthoquinone, is a substance found in all plant chloroplasts. It is, therefore, interesting to know whether it has any influence upon the metabolism of plants. Experiments made with the phytol-free derivatives like 2-methyl-1,4-naphthoquinone or the corresponding 3-oxy compound, phthiocol, gave the following results. These substances accelerate the respiration of Chlorella or Scenedesmus in a way similar to the action of the dinitrophenols. They inhibit photosynthesis and the compensation of respiration in the light strongly like hydroxylamine. In Scenedesmus they hinder the adaptation to the anaerobic utilization of hydrogen. If given after adaptation in amounts sufficient to stop photosynthesis they do not prevent photoreduction but rather stabilize this reaction against reversion. Their presence destroys the coupling between the reduction of carbon dioxide in the dark and the oxyhydrogen reaction in adapted algae. One can expect, therefore, that the natural vitamin K present in plants in concentrations of about 10^–3 M takes part in some metabolic reaction as a catalyst or regulator.     

The temperature dependence of the stimulation of photosynthesis by elevated carbon dioxide in wheat and barley

The temperature dependencies of the solubility of carbon dioxide and oxygen in water and the temperature dependency of the kinetic characteristics of the ribulose-1,5 bisphosphate carboxylase/oxygenase (Rubisco) enzyme result in the short-term stimulation of photosynthesis with a doubling of carbon dioxide from 350 to 700 mol mol^-1 usually decreasing from about 90% at 30C to about 25% at 10C at high photon flux. In field-grown wheat and barley, the expected values at 30°C were observed, but also values as high as 60% at 10°C. The much larger than expected stimulation at cool temperatures in these species also occurred in plants grown at 15°C, but not at 23°C in controlled environment chambers. Gas exchange analysis indicated that an unusually high diffusive limitation was not an explanation for the large response. Assessment of the apparent in vivo specificity of Rubisco by determining the carbon dioxide concentration at which carboxylation equalled carbon dioxide release from oxygenation, indicated that growth at low temperatures altered the apparent enzyme specificity in these species compared to these species grown at the warmer temperature. Inserting the observed specificities into a biochemical model of photosynthesis indicated that altered Rubisco specificity was consistent with the observed rates of assimilation. Whether altered apparent Rubisco specificity is caused by altered stoichiometry of photorespiration or an actual change in enzyme specificity, the results indicate that the temperature dependence of the stimulation of photosynthesis by elevated carbon dioxide may vary greatly with species and with prior exposure to low temperature.Keywords: Barley, carbon dioxide, photosynthesis, temperature, wheat.      

Carbon assimilation by different developmental stages of Laminaria saccharina

Various stages of the life cycle of the marine brown alga Laminaria saccharina (L.) Lamour. (Laminariales, Phaeophyta) including male and female gametophytes, female gametes, zygotes and young sporophytes of different age were investigated for their potentials of carbon dioxide (¹⁴CO₂) fixation. Rates of photosynthesis attain the same order of magnitude in all stages. Photosynthetic ¹⁴CO₂-fixation is accompanied by a substantial light independent carbon assimilation. This is confirmed by rate determinations of the equivalent carboxylating enzymes present in the plants, ribulose-1,5-bisphosphate carboxylase (EC 4.1.1.39) and phosphoenolpyruvate carboxokinase (EC 4.1.1.32) as well as by chromatographic analyses of the appropriate [¹⁴C]-assimilate patterns.Abbreviations RuBP-C ribulose-1,5-bisphosphate carboxylase - PEP-CK phosphoenolpyruvate carboxykinase - PEP phosphoenolpyruvate - PS photosynthesis - DF dark fixation     

Studies on the Biochemistry and Fine Structure of Silica Shell Formation in Diatoms. Photosynthesis and Respiration in Silicon-Starvation Synchrony of Navicula pelliculosa

Rates of photosynthesis, measured by oxygen electrode or by ¹⁴CO₂ fixation, dark respiration and ³²P-phosphate incorporation are reported for the silicon-starvation synchrony of the fresh water diatom Navicula pelliculosa. During late exponential growth the rates were consistent with increase in carbon mass. During silicon starvation, rates of carbon dioxide fixation, oxygen evolution and ³²P incorporation fell, and the saturating light intensity decreased from 27,000 lux to 5000 lux. Reintroduction of silicon led to immediate transients in all parameters studied, followed by a prolonged increase in rate of dark respiration and a gradual increase in apparent photosynthesis. During release of daughter cells, the rates of dark respiration decreased as photosynthesis and ³²P incorporation increased. These results are discussed in relation to effects of silicon on the energy metabolism of the diatom.     

Temporal expression of effects of varying nitrogen supply on canopy growth, photosynthesis and fruit production for Actinidia deliciosa vines in the field

The effects of varying nitrogen supply on canopy leaf area, response of leaf net photosynthesis (A_n) to quantum flux density (Q), and fruit yields of kiwifruit vines (Actinidia deliciosa var. deliciosa) were examined in a two-year field experiment. Vines were grown with 0, 250 or 750 kg N ha^?1 year^?1. The responses to nitrogen supply were compared with responses to shade, to examine the impact of reduced carbon assimilation on canopy leaf area and fruit yields. Nitrogen supply did not affect significantly any of the measured variables during the first season of the experiment. In the second season, canopy leaf area was reduced significantly where nitrogen supply was limited. The quantum efficiency of photosynthesis (φ_q) increased from 0. 03 mol CO₂ mol^?1 Q soon after leaf emergence to more than 0. 05 mol CO₂ mol^?1 Q during the middle of the growing season. The quantum saturated rate of A_n (A_sat) also increased during the season, from 7–10 μmol CO₂ m^?2 s^?1 soon after leaf emergence, to 15–20 (μmol CO₂ m^?2 s^?1 during the middle of the growing season. φ_q and A_sat increased significantly with nitrogen supply at all measurement times during the second season. For vines with high nitrogen, fruit yields in both seasons were similar, averaging 3. 05 kg m^?2. Fruit yields in the second season were reduced significantly where nitrogen supply was limited, due to reduced fruit numbers. The relative effects of reduced leaf area and reduced leaf photosynthesis for carbon assimilation by nitrogen deficient vines were examined using a mathematical model of canopy photosynthesis for kiwifruit vines. Simulations of canopy photosynthesis indicated that effects on leaf area and on leaf photosynthesis were of similar importance in the overall effects of nitrogen deficiency on carbon assimilation. The effects of nitrogen supply on fruit numbers (i. e. flower development) preceded the measured effects on carbon assimilation, indicating that the nitrogen supply affected carbon partitioning to reserves in the first season.     

Demonstration of Both a Photosynthetic and a Nonphotosynthetic CO(2) Requirement for NH(4) Assimilation in the Green Alga Selenastrum minutum

Nitrogen-limited and nitrogen-sufficient cell cultures of Selenastrum minutum (Naeg.) Collins (Chlorophyta) were used to investigate the dependence of NH₄⁺ assimilation on exogenous CO₂. N-sufficient cells were only able to assimilate NH₄⁺ maximally in the presence of CO₂ and light. Inhibition of photosynthesis with 3-(3,4-dichlorophenyl)-1,1-dimethylurea, diuron also inhibited NH₄⁺ assimilation. These results indicate that NH₄⁺ assimilation by N-sufficient cells exhibited a strict requirement for photosynthetic CO₂ fixation. N-limited cells assimilated NH₄⁺ both in the dark and in the light in the presence of 3-(3,4-dichlorophenyl)-1,1-dimethylurea, diuron, indicating that photosynthetic CO₂ fixation was not required for NH₄⁺ assimilation. Using CO₂ removal techniques reported previously in the literature, we were unable to demonstrate CO₂-dependent NH₄⁺ assimilation in N-limited cells. However, employing more stringent CO₂ removal techniques we were able to show a CO₂ dependence of NH₄⁺ assimilation in both the light and dark, which was independent of photosynthesis. The results indicate two independent CO₂ requirements for NH₄⁺ assimilation. The first is as a substrate for photosynthetic CO₂ fixation, whereas the second is a nonphoto-synthetic requirement, presumably as a substrate for the anaplerotic reaction catalyzed by phosphoenolpyruvate carboxylase.     

Mechanism of Photosynthesis Decrease by Verticillium dahliae in Potato

Young, visually symptomless leaves from potato (Solanum tuberosum) plants infected with Verticillium dahliae exhibited reduced carbon assimilation rate, stomatal conductance, and intercellular CO₂, but no increase in dark respiration, no change in the relationship between carbon assimilation rate versus intercellular CO₂, and no change in light use efficiency when intercellular CO₂ was held constant. Therefore, the initial decrease in photosynthesis caused by V. dahliae was caused by stomatal closure. Errors in the intercellular CO₂ calculation caused by uneven distribution of carbon assimilation rate across the leaf were tested by ¹⁴CO₂ autoradiography. Patchiness was found at a low frequency. Low stomatal conductance was correlated with low leaf water potentials. Infection did not affect leaf osmotic potentials.     

Effects of Metabolic Inhibitors on the Calcification of a Freshwater Green Alga, Gloeotaenium loitlesbergarianum Hansgirg. 1. Effects of Some Photosynthetic and Respiratory Inhibitors

The effect of a number of metabolic inhibitors on the calcificationof Gloeotaenium loitlesbergarianum Hansgirg, a freshwater greenalga, was studied. The inhibitors used were methylamine, trimethylamine,mercuric chloride, imidazole, fluoride, arsenate, atrazine,DCMU and dinitrophenol. The effects of these inhibitors showthat transport, or stimulation of respiratory carbon dioxideevolution inhibits calcification. Calcification in Gloeotaeniumis, at least partly, due to a local rise in pH as a result ofphotosynthetic carbon dioxide assimilation. There is also someevidence that, apart from its role in carbon dioxide assimilation,photosynthesis supplies the additional energy needed for calcification. Calcification, Gloeotaenium loitlesbergarianum Hansgirg, green algae, Chlorophyceae, metabolic inhibitors, photosynthesis, respiration     

Short-Term Metabolite Changes during Transient Ammonium Assimilation by the N-Limited Green Alga Selenastrum minutum

In this study, we measured the total pool sizes of key cellular metabolites from nitrogen-limited cells of Selenastrum minutum before and during ammonium assimilation in the light. This was carried out to identify the sites at which N assimilation is acting to regulate carbon metabolism. Over 120 seconds following NH₄⁺ addition we found that: (a) N accumulated in glutamine while glutamate and α-ketoglutarate levels fell; (b) ATP levels declined within 5 seconds and recovered within 30 seconds of NH₄⁺ addition; (c) ratios of pyruvate/phosphoenolpyruvate, malate/phosphoenolpyruvate, Glc-1-P/Glc-6-P and Fru-1,6-bisphosphate/Fru-6-P increased; and (d) as previously seen, photosynthetic carbon fixation was inhibited. Further, we monitored starch degradation during N assimilation over a longer time course and found that starch breakdown occurred at a rate of about 110 micromoles glucose per milligram chlorophyll per hour. The results are consistent with N assimilation occurring through glutamine synthetase/glutamate synthase at the expense of carbon previously stored as starch. They also indicate that regulation of several enzymes is involved in the shift in metabolism from photosynthetic carbon assimilation to carbohydrate oxidation during N assimilation. It seems likely that pyruvate kinase, phosphoenolpyruvate carboxylase, and starch degradation are all activated, whereas key Calvin cycle enzyme(s) are inactivated within seconds of NH₄⁺ addition to N-limited S. minutum cells. The rapid changes in glutamate and triose phosphate, recently shown to be regulators of cytosolic pyruvate kinase, are consistent with them contributing to the short-term activation of this enzyme.     

Stomatal and non-stomatal limitations to carbon assimilation: an evaluation of the path-dependent method

Abstract Environmental stresses can decrease photosynthesis by a direct effect on photosynthetic capacity of the mesophyll or by a CO₂ limitation resulting from stomatal closure. In the present study, a ‘path-dependent method’ (Jones, 1985) for the partitioning of a stress-related decline in assimilation rate between non-stomatal and stomatal factors was evaluated, using light quality as a ‘stress’. Kinetic data on assimilation rate and conductance of Phragmipedium longifolium following a change in light quality from 95 μmol m^?2s^?1 white light to 95 μmol m^?2s^?1 red light failed to generate a smooth response curve for conductance. Partitioning of limitations on assimilation by a path-dependent method that utilizes the actual trajectories of conductance and assimilation was therefore not feasible. A simplified path-dependent method (Jones, 1985) which assumes that either mesophyll cells or guard cells respond first to a stress was applied to steady-state measurements of assimilation and conductance under red and white illumination. Either 5% or 23% of the observed reduction in assimilation rate under white light was attributable to stomatal factors, depending on whether the ‘stomatal first’ or the ‘mesophyll first’ path was assumed. In the absence of additional information indicating the appropriate choice of path, arbitrary choice may therefore lead to widely divergent estimates, and potentially erroneous conclusions. An alternative approach to the evaluation of the importance to carbon assimilation of stomatal and non-stomatal factors is suggested.     

Microbial processes at the aerobic-anaerobic interface in the deep-water zone of the black sea

Chemical and key microbiological processes (assimilation of carbon dioxide, oxidation and formation of methane, and sulfate reduction) occurring at the aerobic-anaerobic interface in the deep-water zone of the Black Sea were investigated. Measurements were taken at depths from 90 to 300 m at intervals of 5–10 m. The integral rate of the dark assimilation of carbon dioxide varied from 120 to 207 mg C/(m² day) with a maximum at the boundary of cyclonic currents. The organic matter (OM) formed from methane comprised less than 5% of the OM formed from carbon dioxide. A comparison between the rates of methane oxidation and methane production suggests that methane that is oxidized at depths from 100 to 300 m was formed in deeper water horizons. The maximum rate of sulfate reduction (1230 mg S/(m₂ day)) was observed in the western halistatic region, and the minimum rate (490 mg S/(m₂ day)), in the eastern halistatic region. The average rate of hydrogen sulfide production measured at three deep-sea stations amounted to 755 mg S/(m₂ day), or 276 g S/(m₂ year).